Arthropod touch reception: spider hair sensilla as rapid touch detectors

Wandering spiders like Cupiennius salei are densely covered by tactile hairs. In darkness Cupiennius uses its front legs as tactile feelers. We selected easily identifiable hairs on the tarsus and metatarsus which are stimulated during this behavior to study tactile hair properties. Both the mechanical and electrophysiological hair properties are largely independent of the direction of hair displacement. Restoring torques measure 10(-9) to 10(-8) Nm. The torsional restoring constant S changes non-linearly with deflection angle. It is of the order of 10(-8) Nm/rad, which is about 10,000 times larger than for trichobothria. Angular thresholds for the generation of action potentials are ca.1 degrees. Electrophysiology reveals a slow and a fast sensory cell, differing in adaptation time. Both cells are movement detectors mainly responding to the dynamic phase (velocity) of a stimulus. When applying behaviorally relevant stimulus velocities (up to 11 cm s(-1)) threshold deflection for the elicitation of action potentials and maximum response frequency are reached as early as 1.2 ms after stimulus onset and followed by a rapid decline of impulse frequency. Obviously these hairs inform the spider on the mere presence of a stimulus but not on details of its time-course and spatial orientation.     

Arthropod touch reception: structure and mechanics of the basal part of a spider tactile hair

Our previous studies on spider tactile hairs concentrated on the mechanical behavior of the hair shaft and the electrophysiological properties of the sensory cells. Here we focus on the structure and mechanical properties of the coupling of the hair shaft and the sensory terminals. 1. The functional design of the coupling provides for a combination of high sensitivity and protection against mechanical damage and overstimulation. The dendritic sheath is not directly coupled to the hair shaft. Rather, there is terminal connecting material between the dendrites and the hair shaft. 2. The hair shaft forms a first-order lever. Its acentric axis of rotation is located ca.3.5 m from its inner end. Displacement of the hair tip is scaled down by a factor of ca.750:1, not even considering the outer hair shafts bending. 3. At threshold the dendrite sheath displacement is ca. 0.05 m by forces in the order of 0.4–4×10^–6 N. 4. The hair shaft bends within the socket even before contacting it. The elasticities representing its suspension and bending in the socket can be described quantitatively by measuring the hairs restoring moments (range: 10^–9 Nm) and bending at different degrees of deflection.     

Mechanosensory afferents innervating the swimmerets of the lobster

Feathered hair sensilla fringe both rami of the lobster (Homarus americanus) swimmeret. The sensory response to hair displacement was characterized by recording afferent impulses extracellularly from the swimmeret sensory nerve while deflecting sensilla with a rigidly-coupled probe or controlled water movements. Two populations of hairs were observed: "distal" hairs localized to the distal 1/3 of each ramus and "proximal" hairs near its base. Distal hairs are not innervated by a mechanosensory neuron but instead act as levers producing strain within adjacent cuticle capable of activating a nearby hypodermal mechanoreceptor. Hair deflections of 25 degrees or more are required to evoke an afferent response and this response is dependent on hair deflection direction. The frequency and duration of the afferent discharge evoked are determined by the velocity of hair displacement. Each proximal hair is innervated by a single mechanosensory neuron responding phasically to hair deflections as small as 0.2 degrees in amplitude. Deflection at frequencies up to 5 Hz elicits a single action potential for each hair movement; at higher frequencies many deflections fail to evoke an afferent response. These sensilla, which are mechanically coupled, may be activated by the turbulent flow of water produced by the swimmerets during their characteristic beating movements.     

The Advantages of a Tapered Whisker

The role of facial vibrissae (whiskers) in the behavior of terrestrial mammals is principally as a supplement or substitute for short-distance vision. Each whisker in the array functions as a mechanical transducer, conveying forces applied along the shaft to mechanoreceptors in the follicle at the whisker base. Subsequent processing of mechanoreceptor output in the trigeminal nucleus and somatosensory cortex allows high accuracy discriminations of object distance, direction, and surface texture. The whiskers of terrestrial mammals are tapered and approximately circular in cross section. We characterize the taper of whiskers in nine mammal species, measure the mechanical deflection of isolated felid whiskers, and discuss the mechanics of a single whisker under static and oscillatory deflections. We argue that a tapered whisker provides some advantages for tactile perception (as compared to a hypothetical untapered whisker), and that this may explain why the taper has been preserved during the evolution of terrestrial mammals.     

Measurement of the force produced by an intact bull sperm flagellum in isometric arrest and estimation of the dynein stall force

The force generated by a detergent-extracted reactivated bull sperm flagellum during an isometric stall was measured with a force-calibrated glass microprobe. The average isometric stall force from 48 individual measurements was 2.5 +/- 0.7 x 10(-5) dyne (2.5 +/- 0.7 x 10(-10) N). The force measurements were obtained by positioning a calibrated microprobe in the beat path of sperm cells that were stuck by their heads to a glass microscope slide. The average position of the contact point of the flagellum with the probe was 15 microm from the head-tail junction. This average lever arm length multiplied by the measured force yields an estimate of the active bending moment (torque) of 3.9 x 10(-8) dyne x cm (3.9 x 10(-15) N x m). The force was sustained and was for the most part uniform, despite the fact that the flagellum beyond the point of contact with the probe usually continued beating. It appears that the dynein motors in the basal portion of the flagellum continue to pull in an isometric stall for as long as the motion of the flagellum is blocked. If dynein motors in the flagellum distal to the contact point with the probe were contributing force to the displacement of the probe, then the flagellar segment immediately past the point of contact would have to show a net curvature in the direction of the probe displacement. No such curvature bias was observed in the R-bend arrests, and only a small positive curvature bias was measured in the P-bend arrests. Our analysis of the data suggests that more than 90% of the sustained force component is generated by the part of the flagellum between the probe and the flagellar base. Based on this premise, the isometric stall force per dynein head is estimated to be 5.0 x 10(-7) dyne (5 pN). This equals approximately 1.0 x 10(-6) dyne (10 pN) per intact dynein arm. These values are close to the isometric stall force of isolated dynein. This suggests that all of the dynein heads between the base and the probe, on the active side of the axoneme, are contributing to the force exerted against the probe.     

Tactile localisation: the function of active antennal movements in the crayfishCherax destructor

Video recordings and single frame analysis were used to study the function of the second antennae of crayfish (Cherax destructor) as a sensory system in freely behaving animals. Walking crayfish move their antennae back and forth through horizontal angles of 100 degrees and more, relative to the body long axis. At rest, animals tend to hold their antennae at angular positions between 20 and 50 degrees. Movements of the two antennae are largely independent of each other. Before and during a turn of the body the ipsilateral antenna is moved into the direction of the turn. Solid objects are explored by repeatedly moving the antennae towards and across them. Both seeing and blinded crayfish can locate stationary objects following antennal contact. On antennal contact with a small novel object, a moving animal withdraws its antenna and attacks the object. When the antenna of a blinded crayfish is lightly touched with a brush the animal turns and attacks the point of stimulation. The direction taken and the distance covered during an attack can be correlated with: the angle at which the antenna is held at the moment of contact and the distance along the antennal flagellum at which the stimulus is applied. From behavioural evidence we conclude that crayfish use information about the angular position of their antennae and about the position of stimulated mechanoreceptors along the antennal flagellum to locate objects in their environment. We suggest ways in which an active tactile system-like the crayfish's antennae--could supply animals with information about the three-dimensional layout of their environment.     

Structural scaling and functional design of the cercal wind-receptor hairs of cricket

We have estimated the intrinsic mechanical parameters of cricket cercal wind-receptor hairs. The hairs were modeled as an inverted pendulum, and mechanical parameters of the equation of motion were determined from data given by a systematic measurement of mobility by the least-square error method. The theoretical torque which turns the hair shaft is given by the drag force due to the moving air. The drag force is given by the method of Stokes' mechanical impedance of an oscillating cylinder in viscous fluid. The effect of the boundary layer in which air is stagnating on the substrate surface is also taken into account. The moment of inertia of a hair shaft shows a clear length dependency to the power of 4.32 of the hair length. The torsional resistance within the hair base and the stiffness of hair-supporting spring also show clear length dependencies to the power of 2.77 and 1.67, respectively. The torsional resistance within the hair base is so large that the hair is a strongly damped non-oscillatory second-order system. The large resistance within the hair base represents an efficient energy absorption by the sensory cell. The resistance seems to match with the source impedance, i.e., the frictional resistance at the site of air-hair contact. The impedance matching provides the condition of maximum power transmission from the moving air to the sensory cell. Structural scaling is discussed in relation to the functional scaling of the frequency-range fractionation of the mechanical filter array with a common biological design. Accepted: 24 February 1998     

The morphology of spider sensilla. I. Mechanoreceptors

The common tactile hair sensilla of spider tarsi were studied in web spiders (Araneus) and ground spiders (Lycosa, Dugesiella) using scanning and transmission electron microscopy. All of these sensilla are innervated by three bipolar neurons whose dendrites end proximally at the sensillum base. Each dendritic terminal exhibits a tubular body, a dense array of microtubules typical for mechanoreceptive sensilla. A dendritic sheath encloses the outer dendritic segments and connects the dendritic terminals to cuticular components of the hair sensillum in three different ways: (1) A distal extension of the dendritic sheath connects to the midline of the hair base; (2) A forked arrangement of cuticular (?) strands attaches on both lateral sides of the hair base, and (3) The socket cuticle directly contacts a part of the dendritic sheath. The latter connection provides a fixed position for the three dendritic terminals and any movement of the hair shaft could be transmitted via connections (I) and (2). The triple innervation strongly suggests a directional sensitivity of these sensilla.Structural comparison between arachnid and insect mechanoreceptive sensilla indicates that tactile hair sensilla in Arachnida are multi-innervated whereas the corresponding reccptors in Insecta are singly innervated.     

Mechanical properties of the lateral cortex of mammalian auditory outer hair cells.

Mammalian auditory outer hair cells generate high-frequency mechanical forces that enhance sound-induced displacements of the basilar membrane within the inner ear. It has been proposed that the resulting cell deformation is directed along the longitudinal axis of the cell by the cortical cytoskeleton. We have tested this proposal by making direct mechanical measurements on outer hair cells. The resultant stiffness modulus along the axis of whole dissociated cells was 3 x 10(-3) N/m, consistent with previously published values. The resultant axial and circumferential stiffness moduli for the cortical lattice were 5 x 10(-4) N/m and 3 x 10(-3) N/m, respectively. Thus the cortical lattice is a highly orthotropic structure. Its axial stiffness is small compared with that of the intact cell, but its circumferential stiffness is within the same order of magnitude. These measurements support the theory that the cortical cytoskeleton directs electrically driven length changes along the longitudinal axis of the cell. The Young's modulus of the circumferential filamentous components of the lattice were calculated to be 1 x 10(7) N/m2. The axial cross-links, believed to be a form of spectrin, were calculated to have a Young's modulus of 3 x 10(6) N/m2. Based on the measured values for the lattice and intact cell cortex, an estimate for the resultant stiffness modulus of the plasma membrane was estimated to be on the order of 10(-3) N/m. Thus, the plasma membrane appears to be relatively stiff and may be the dominant contributor to the axial stiffness of the intact cell.     

Characteristics of serial cross-sections of hairs of the Japanese monkey (Macaca fuscata fuscata)

The characteristics of serial cross-sections of hairs collected from an adult male Japanese monkey were investigated. Cross-sections were made of five to eight pieces per hair. The shapes of the cross-sections were elliptical or rounded on the whole. The fibre indices of the sections ranged from 83 to 100. In particular, those of proximal (basal) sections were close to 100. The hair diameter was 86.4 μ at maximum and 27.2 μ at minimum. A tendency was observed for the longer hairs to have thicker diameters. The changes in thickness along the fibre shaft were slightly different in relation to hair length. The thickest point was at around the middle of the fibre in the intermediate hair, somewhat towards the top of the central part in the long hair, and somewhat towards the base in the short hair. The hair of the Japanese monkey, however, was considered to be scanty in changes along the fibre shaft in comparison with many other animals. Medullae could scarcely be seen in the short hair and in the terminal and proximal sections of all hairs. Their shapes in cross-section were not uniform and rough at the margins. The fibre-medulla indices were generally less than 30 and smaller than those of many other mammals. Pigmentary granules were observed in all sections examined. The granules were black-grey in sections of the black-grey coloured part and yellow in the yellowish sections. They were dense in distal sections and scarce in sections close to the base. The cross-sectional appearance of the thickest part of the long hair was considered to be useful for hair identification, since it was good in pigmentation and medullation and relatively small fibre index.     

Reaction of single cercal mechanoreceptors of the cricket, Gryllus bimaculatus, to mechanical stimulation

Naked tungsten microelectrodes were introduced through the chitinous wall close to the cell body of the receptor cells of trichoid hair sensillae, and responses to airpuffs, and rectangular, and trapezoidal displacements of the hair were recorded. Receptors of dorsal zone are activated during lateral deflection, those of ventral zone--during medial deflection and receptors of medial and lateral zones--during deflection to the cercal base. Sensitivity of receptors to the air-puffs is a function of hair length, the largest hairs being most sensitive. During trapezoidal displacement of the hair with different velocities of the slope, discharge frequency of the dynamic response is a function of velocity and angle in the range of angles up to 3-5 degrees (fig. 1). Discharge frequency of the stationary phase (corresponding to the plateau of the stimulus) is mainly a function of velocity in the range up to 6 degrees (fig. 2). The presence of sensillae with different hair length, and hence sensitivity, and definite directionality of receptors in different hair length, and hence sensitivity, and definite directionality of receptors in different zones may provide a basis for amplitude, velocity and direction discrimination of air-puffs or low-frequency mechanical stimulation by the cercal system of crickets.     

Mechanosensory control of antennal movement by the scapal hair plate in the American cockroach

The voluntary movement of antennae of blinded cockroaches was examined in the tethered-walking condition. An object of metal plate was presented to a tip of a single antenna in order to induce tactile orientation behavior. Horizontal movements of the antenna before and during the object presentation were analyzed both before and after ablation of a mechanosensory organ, the scapal hair plate (S-HP), at the base of antenna. The resting antennal position shifted outwardly by about 20 degrees after the S-HP ablation. Spontaneous antennal movements in ablated animals became stiff and wider ranged. The tactile object was set at two different horizontal positions, 45 degrees and 90 degrees clockwise to the head, for the right side antenna. The number of contacts in a constant test period was significantly decreased in the tests at 45 degrees after ablation. Trajectories of antennal movements before and after contacts were categorized into four patterns. In the case that an antenna made contact with the object during its abduction (outward) movement, it then passed the object outwardly or withdrew inwardly. These were termed "outward-pass (O-P)" and "outward-withdrawal (O-W)" patterns, respectively. Similarly, contacts during the adduction (inward) movement were divided into "inward-pass (I-P)" or "inward-withdrawal (I-W)" pattern. Significant effects of the S-HP ablation appeared in the tests at 90 degrees : the I-P pattern mostly disappeared and was replaced by the I-W pattern. The results strongly suggest that the S-HP has crucial roles for controlling both spontaneous and stimulated movements of the cockroach antenna.     

Responses of cutaneous mechanoreceptors within fingerpad to stimulus information for tactile softness sensation of materials

Softness sensation is one of primitive tactile textures. While the psychophysical characteristics of softness sensation have been thoroughly studied, it is lack of a deep understanding of the underlying neuromechanical principles. On the stimulus–response processes of human fingerpad touching fabrics and the physiological properties of slowly adapting type I (SAIs) cutaneous mechanoreceptors within fingerpad, a fabric-skin-receptor coupling model was built and validated. By the fabric-skin-receptor model a series of numerical experiments was conducted, and how the evoked neural responses of cutaneous mechanoreceptors change with the composite compliance of both fingerpad skin and the materials in contact was investigated. The results indicated that the evoked neural responses of populations of cutaneous mechanoreceptors by the physical stimulus from fabrics were nearly proportional to the perceived softness magnitude, and nonlinearly increased and then decreased with the effective elastic modulus of fabrics or the relative elastic modulus of fabrics to soft tissues within fingerpad, where the nonlinear inflection point depended on the touching force level. Therefore, it concluded that the tactile judgment of the physical information for softness sensation of objects was an encoding of neural responses of populations of SAIs cutaneous mechanoreceptors, and the physical information depended on the mechanical interaction of fingerpad and objects in contact.     

Chemosensitive hairs in spiders

Spiders possess curved, blunt-tipped hairs on all legs and palps, which differ in many details from the straight, sharp-pointed, tactile hairs: (1) the blunt tip is open to the outside, which can be demonstrated by high resolution microscopy and by the penetration of dyes; (2) the hair shaft has a double lumen which consists of a circular (tube) and a crescent shaped lumen; (3) this hair is innervated by two to three bipolar neurons whose dendrites enter the small tube, where they arborize into 16–20 branches. Multiple innervation and an open tip give strong evidence for a chemoreceptive function. Concluding from their position and distribution on the distal leg parts, a contact chemoreception is tentatively proposed. This interpretation is supported by the close structural analogy to the known contact chemoreceptors in insects. Observation of behavior indicates the importance of a contact chemoreceptor on spider legs. Other possible chemoreceptors in spiders which have been described previously by other authors are discussed.     

Physical and physiological properties of the crayfish antennal flagellum.

(1) Mechanoreceptors in the crayfish antennae are divided into four functional categories: vibration (13%), bidirectional displacement (19%), unidirectional displacement (45%), and position (23%) receptors. The distribution of receptors along the length of the flagellum follows a logarithmic progression, decreasing from about 40% at the base to less than 5% at the tip. (2) Vibratory stimulation of the antennae was found to induce a traveling wave. Because of an impedance gradient along the length of the flagellum, the traveling wave moves most efficiently from base to tip. The wave was observed to travel at an average velocity of 6.0 m/sec. (3) Large deflections of the tip are not uniformly transferred to the base, but decrease logarithmically. This due to the existence of the impedance gradient. (4) Receptor output probability was found to be greatest when low frequency/high intensity stimulation was applied to the flagellar base. (5) Characteristics of large (2 cm) posterior-going deflections of the flagellar tip that are effective in producing response differences are displacement: (a) amplitude, (b) velocity, and (c) acceleration.     

Design and characterization of asparagine- and lysine-containing alanine-based helical peptides that bind selectively to A.T base pairs of oligonucleotides immobilized on a 27 mhz quartz crystal microbalance

We have systematically designed and synthesized six kinds of 16-17 mer alanine-based peptides containing four to six lysine (K) and one to four asparagine (N) residues to achieve the selective binding to A.T base pairs of DNA duplexes. The position and number of K and N residues were changed in the helical structure according to common features of the DNA-binding proteins, in which K and N residues are expected to interact electrostatically with phosphate groups and to interact with A.T base pairs by hydrogen bonding, respectively. The time courses of binding of these peptides to dA(30).dT(30) and dG(30).dC(30) duplexes immobilized on a 27 MHz quartz crystal microbalance (QCM) were studied in 10 mM phosphate buffer (pH 7.5) and 40 mM NaCl at 10 degrees C. The maximum binding amounts (Deltam(max)) on a nanogram scale and binding constants (K(a)) could be obtained from the frequency decrease (mass increase) of the oligonucleotide-immobilized QCM. The conformation changes of the peptides upon binding to DNAs were monitored by circular dichroism (CD) spectroscopy. The four properly arranged N residues in the six-cationic K peptide, K6N4(d), resulted in a 5-fold higher affinity for A.T base pairs (K(a) = 5.9 x 10(5) M(-1)) than for G.C base pairs (K(a) = 1.2 x 10(5) M(-1)), and alpha-helices were clearly promoted by the binding to A.T base pairs from CD spectral changes.     

Hair analysis of histamine after fluorescence labeling by column-switching reversed-phase liquid chromatography with electrospray ionization mass spectrometry and application to human hair

Sensitive determination of histamine (HA) in hair was carried out by column-switching reversed-phase high-performance liquid chromatography coupled with electrospray ionization mass spectrometry (HPLC-ESI-MS). HA was labeled with excess amounts of 4-(N,N-dimethylaminosulfonyl)-7-fluoro-2,1,3-benzoxadiazole (DBD-F) at 60 degrees C for 30 min in a mixture of 0.1 M borax (pH 9.3) and acetonitrile (CH(3)CN). The resulting DBD-HA derivative was roughly separated by a Mightysil RP-18 GP (100 x 2mm i.d., 3 microm) with an acidic mobile phase containing 0.1% trifluoroacetic acid. DBD-HA in the fraction flowing due to a position change in the six-port column-switching valve was then completely separated by a Wakopak Navi C30 (150 x 2mm i.d., 5 microm) with 20 mM AcONH(4)-CH(3)CN (8:2). The mass spectrometer was operated in the selected reaction monitoring (SRM) mode for the product ion (m/z 292) obtained from MS-MS measurement using the protonated molecular ion [M+H](+) (m/z 337) as the precursor ion. Good linearity was achieved from the calibration curve obtained by plotting peak area ratios of the internal standard (HA-d(4)) against the injected amounts of HA (1.66-16.6 pmol, r(2)=0.999). The coefficients of variation, at 1.66- and 16.6-pmol injections, were 5.6 and 3.7%, respectively (n=6). Furthermore, the detection limit was 0.167 pmol. The efficiency of the recommended procedure was identified from the determination in the rat hair root after intraperitoneal administration of HA. The proposed method was applied to HA determination in the hair shaft of Dark Agouti rats and healthy volunteers. The variations in the concentrations in 1mg of hair shaft were 0.80-1.84 pmol (mean+/-SD=1.33+/-0.33, n=12) in rats and 0.94-72.3 pmol (17.2+/-21.5, n=16) in humans. The determination of HA in the plasma of rats and humans was also performed successfully by this method. Because the proposed method provides good precision and trace detection of HA in hair, the analytical technique seems to be applicable for the determination of various biogenic amines in hair.     

Tibiofemoral joint contact force in deep knee flexion and its consideration in knee osteoarthritis and joint replacement

The aim of the study was to estimate the tibiofemoral joint force in deep flexion to consider how the mechanical load affects the knee. We hypothesize that the joint force should not become sufficiently large to damage the joint under normal contact area, but should become deleterious to the joint under the limited contact area. Sixteen healthy knees were analyzed using a motion capture system, a force plate, a surface electromyography, and a knee model, and then tibiofemoral joint contact forces were calculated. Also, a contact stress simulation using the contact areas from the literature was performed. The peak joint contact forces (M +/- SD) were 4566 +/- 1932 N at 140 degrees in rising from full squat and 4479 +/- 1478 N at 90 degrees in rising from kneeling. Under normal contact area, the tibiofemoral contact stresses in deep flexion were less than 5 MPa and did not exceed the stress to damage the cartilage. The contact stress simulation suggests that knee prosthesis having the contact area smaller than 200 mm2 may be problematic since the contact stress in deep flexion would become larger than 21 MPa, and it would lead damage or wear of the polyethylene.     

Effects of fusion between tactile and proprioceptive inputs on tactile perception

Tactile perception is typically considered the result of cortical interpretation of afferent signals from a network of mechanical sensors underneath the skin. Yet, tactile illusion studies suggest that tactile perception can be elicited without afferent signals from mechanoceptors. Therefore, the extent that tactile perception arises from isomorphic mapping of tactile afferents onto the somatosensory cortex remains controversial. We tested whether isomorphic mapping of tactile afferent fibers onto the cortex leads directly to tactile perception by examining whether it is independent from proprioceptive input by evaluating the impact of different hand postures on the perception of a tactile illusion across fingertips. Using the Cutaneous Rabbit Effect, a well studied illusion evoking the perception that a stimulus occurs at a location where none has been delivered, we found that hand posture has a significant effect on the perception of the illusion across the fingertips. This finding emphasizes that tactile perception arises from integration of perceived mechanical and proprioceptive input and not purely from tactile interaction with the external environment.     

The effect of axial compression and distraction on cervical facet mechanics during anterior shear,flexion, axial rotation,and lateral bending motions

The subaxial cervical facets are important load-bearing structures, yet little is known about their mechanical response during physiological or traumatic intervertebral motion. Facet loading likely increases when intervertebral motions are superimposed with axial compression forces, increasing the risk of facet fracture. The aim of this study was to measure the mechanical response of the facets when intervertebral axial compression or distraction is superimposed on constrained, non-destructive shear, bending and rotation motions. Twelve C6/C7 motion segments (70 ± 13 yr, nine male) were subjected to constrained quasi-static anterior shear (1 mm), axial rotation (4°), flexion (10°), and lateral bending (5°) motions. Each motion was superimposed with three axial conditions: (1) 50 N compression; (2) 300 N compression (simulating neck muscle contraction); and, (3) 2.5 mm distraction. Angular deflections, and principal and shear surface strains, of the bilateral C6 inferior facets were calculated from motion-capture data and rosette strain gauges, respectively. Linear mixed-effects models (α = 0.05) assessed the effect of axial condition. Minimum principal and maximum shear strains were largest in the compressed condition for all motions except for maximum principal strains during axial rotation. For right axial rotation, maximum principal strains were larger for the contralateral facets, and minimum principal strains were larger for the left facets, regardless of axial condition. Sagittal deflections were largest in the compressed conditions during anterior shear and lateral bending motions, when adjusted for facet side.